The two basic phenomena that define the problem of visual attention can be illustrated in a simple example. Consider the arrays shown in each panel of Figure 1. In a typical experiment, before the arrays were presented, subjects would be asked to report letters appearing in one color (targets, here black letters), and to disregard letters in the other color (nontargets, here white letters). The array would then be briefly flashed, and the subjects, without any opportunity for eye movements, would give their report. The display mimics our. usual cluttered visual environment: It contains one or more objects that are relevant to current behavior, along with others that are irrelevant. The first basic phenomenon is limited capacity for processing information. At any given time, only a small amount of the information available on the retina can be processed and used in the control of behavior. Subjectively, giving attention to any one target leaves less available for others. In Figure 1, the probability of reporting the target letter N is much lower with two accompa­ nying targets (Figure la) than with none (Figure Ib). The second basic phenomenon is selectivity-the ability to filter out un­ wanted information. Subjectively, one is aware of attended stimuli and largely unaware of unattended ones. Correspondingly, accuracy in identifying an attended stimulus may be independent of the number of nontargets in a display (Figure la vs Ie) (see Bundesen 1990, Duncan 1980).

Neural Mechanisms of Selective Visual Attention

Sensory information undergoes extensive associative elaboration and attentional modulation as it becomes incorporated into the texture of cognition. This process occurs along a core synaptic hierarchy which includes the primary sensory, upstream unimodal, downstream unimodal, heteromodal, paralimbic and limbic zones of the cerebral cortex. Connections from one zone to another are reciprocal and allow higher synaptic levels to exert a feedback (top-down) influence upon earlier levels of processing. Each cortical area provides a nexus for the convergence of afferents and divergence of efferents. The resultant synaptic organization supports parallel as well as serial processing, and allows each sensory event to initiate multiple cognitive and behavioural outcomes. Upstream sectors of unimodal association areas encode basic features of sensation such as colour, motion, form and pitch. More complex contents of sensory experience such as objects, faces, word-forms, spatial locations and sound sequences become encoded within downstream sectors of unimodal areas by groups of coarsely tuned neurons. The highest synaptic levels of sensory-fugal processing are occupied by heteromodal, paralimbic and limbic cortices, collectively known as transmodal areas. The unique role of these areas is to bind multiple unimodal and other transmodal areas into distributed but integrated multimodal representations. Transmodal areas in the midtemporal cortex, Wernicke's area, the hippocampal-entorhinal complex and the posterior parietal cortex provide critical gateways for transforming perception into recognition, word-forms into meaning, scenes and events into experiences, and spatial locations into targets for exploration. All cognitive processes arise from analogous associative transformations of similar sets of sensory inputs. The differences in the resultant cognitive operation are determined by the anatomical and physiological properties of the transmodal node that acts as the critical gateway for the dominant transformation. Interconnected sets of transmodal nodes provide anatomical and computational epicentres for large-scale neurocognitive networks. In keeping with the principles of selectively distributed processing, each epicentre of a large-scale network displays a relative specialization for a specific behavioural component of its principal neurospychological domain. The destruction of transmodal epicentres causes global impairments such as multimodal anomia, neglect and amnesia, whereas their selective disconnection from relevant unimodal areas elicits modality-specific impairments such as prosopagnosia, pure word blindness and category-specific anomias. The human brain contains at least five anatomically distinct networks. The network for spatial awareness is based on transmodal epicentres in the posterior parietal cortex and the frontal eye fields; the language network on epicentres in Wernicke's and Broca's areas; the explicit memory/emotion network on epicentres in the hippocampal-entorhinal complex and the amygdala; the face-object recognition network on epicentres in the midtemporal and temporopolar cortices; and the working memory-executive function network on epicentres in the lateral prefrontal cortex and perhaps the posterior parietal cortex. Individual sensory modalities give rise to streams of processing directed to transmodal nodes belonging to each of these networks. The fidelity of sensory channels is actively protected through approximately four synaptic levels of sensory-fugal processing. The modality-specific cortices at these four synaptic levels encode the most veridical representations of experience. Attentional, motivational and emotional modulations, including those related to working memory, novelty-seeking and mental imagery, become increasingly more pronounced within downstream components of unimodal areas, where they help to create a highly edited subjective version of the world. (ABSTRACT TRUNCATED)

http://zielinskifam.com/lit/gumc_nsci_papers/Mesulam_98_Brain_cog_review.pdf

From sensation to cognition.

By utilizing new information from both clinical and experimental (lesion, electrophysiological, and gene-activation) studies with animals, the anatomy underlying anterograde amnesia has been reformulated. The distinction between temporal lobe and diencephalic amnesia is of limited value in that a common feature of anterograde amnesia is damage to part of an “extended hippocampal system” comprising the hippocampus, the fornix, the mamillary bodies, and the anterior thalamic nuclei. This view, which can be traced back to Delay and Brion (1969), differs from other recent models in placing critical importance on the efferents from the hippocampus via the fornix to the diencephalon. These are necessary for the encoding and, hence, the effective subsequent recall of episodic memory. An additional feature of this hippocampal–anterior thalamic axis is the presence of projections back from the diencephalon to the temporal cortex and hippocampus that also support episodic memory. In contrast, this hippocampal system is not required for tests of item recognition that primarily tax familiarity judgements. Familiarity judgements reflect an independent process that depends on a distinct system involving the perirhinal cortex of the temporal lobe and the medial dorsal nucleus of the thalamus. In the large majority of amnesic cases both the hippocampal–anterior thalamic and the perirhinal–medial dorsal thalamic systems are compromised, leading to severe deficits in both recall and recognition.

/pdf/episodic-memory-amnesia-and-the-hippocampal-anterior-50qwk8ap9b.pdf

Episodic memory, amnesia, and the hippocampal–anterior thalamic axis.

One of the enduring mysteries of brain function concerns the process of cognitive control. How does complex and seemingly willful behaviour emerge from interactions between millions of neurons? This has long been suspected to depend on the prefrontal cortex--the neocortex at the anterior end of the brain--but now we are beginning to uncover its neural basis. Nearly all intended behaviour is learned and so depends on a cognitive system that can acquire and implement the 'rules of the game' needed to achieve a given goal in a given situation. Studies indicate that the prefrontal cortex is central in this process. It provides an infrastructure for synthesizing a diverse range of information that lays the foundation for the complex forms of behaviour observed in primates.

/pdf/the-prefrontal-cortex-and-cognitive-control-154fw1z9a0.pdf

The prefrontal cortex and cognitive control.

The hallmark of medial temporal lobe amnesia is a loss of episodic memory such that patients fail to remember new events that are set in an autobiographical context (an episode). A further symptom is a loss of recognition memory. The relationship between these two features has recently become contentious. Here, we focus on the central issue in this dispute — the relative contributions of the hippocampus and the perirhinal cortex to recognition memory. A resolution is vital not only for uncovering the neural substrates of these key aspects of memory, but also for understanding the processes disrupted in medial temporal lobe amnesia and the validity of animal models of this syndrome.

Recognition memory: what are the roles of the perirhinal cortex and hippocampus?

The activity of 736 single neurons was recorded from the hippocampal formation (HF), the rhinal cortex (RH), the medial and anterior inferior temporal cortex (TE), or areas TF and TH of the parahippocampal gyrus (PHG) of monkeys during the performance of a delayed matching to sample task. The results indicate differences between the areas in their contributions to sensory processing and memory. Of the neurons, 55% responded to either the first (S1) and/or the second (S2) of the two successively presented visual stimuli. The proportion of responsive neurons and the proportion of neurons that responded selectively on the basis of shape or color (but not size) were significantly higher in areas TE + RH than in HF + PHG. The responses to S1 differed from those to S2 for 18% of the total sample: of these differentially responsive neurons, 66% of the TE + RH neurons responded more strongly to S1 (the sample presentation, allowing stimulus acquisition), whereas 71% of the HF + PHG neurons responded more strongly to S2 (the match/nonmatch comparison, when the behavioral decision could be made). Of 239 TE + RH neurons recorded during the delayed matching task or when objects were shown, 12% displayed evidence of memory for the previous occurrence of stimuli by responding strongly to the first, but significantly less strongly to subsequent presentations of visual stimuli that were novel or had not been seen recently. In contrast, none (0%) of 328 neurons so tested in HF and PHG had a response that declined significantly on stimulus repetition. For six (86%) of seven TE + RH neurons tested, the decrement in response persisted even after distraction by intervening presentations of other stimuli. Further evidence of information storage was found for 7 (33%) of 21 neurons for which responses to the first presentations of unfamiliar objects were significantly greater than to the first presentations of very familiar objects, even though the familiar objects had not been seen for greater than 15 min.

https://www.jneurosci.org/content/jneuro/11/6/1763.full.pdf

The effects of visual stimulation and memory on neurons of the hippocampal formation and the neighboring parahippocampal gyrus and inferior temporal cortex of the primate.

Neuronal evidence that inferomedial temporal cortex is more important than hippocampus in certain processes underlying recognition memory

Hippocampus and medial temporal cortex: neuronal activity related to behavioural responses during the performance of memory tasks by primates.

Neurones recorded from the monkey medial temporal lobe display evidence of memory for the previous occurrence of visual stimuli. In the inferomedial temporal cortex, 26 (20%) of 128 single units responded more strongly to the first presentations of visual stimuli which had not been seen recently than to subsequent presentations. Such declining responses were also found in the basolateral amygdala and ventral putamen. No such responses were found in the hippocampus, dentate gyrus or subicular cortex. These findings suggest that the inferomedial temporal cortex, rather than the hippocampal formation, plays a central role in recognition memory.

F. A. W. Wilson

Papers

The effects of visual stimulation and memory on neurons of the hippocampal formation and the neighboring parahippocampal gyrus and inferior temporal cortex of the primate.

Neuronal evidence that inferomedial temporal cortex is more important than hippocampus in certain processes underlying recognition memory

Hippocampus and medial temporal cortex: neuronal activity related to behavioural responses during the performance of memory tasks by primates.

Neurones of the Medical Temporal Lobes and Recognition Memory